Research Article |
Corresponding author: Xiao-Dan Yu ( yuxd126@126.com ) Academic editor: Zai-Wei Ge
© 2021 Ying Pei, Hong-Bo Guo, Tie-Zhi Liu, Wei-Qiang Qin, Di Zhao, Xiao-Jian Qi, Xiao-Dan Yu.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pei Y, Guo H-B, Liu T-Z, Qin W-Q, Zhao D, Qi X-J, Yu X-D (2021) Three new Melanoleuca species (Agaricales, Basidiomycota) from north-eastern China, supported by morphological and molecular data. MycoKeys 80: 133-148. https://doi.org/10.3897/mycokeys.80.64369
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Three new Melanoleuca species, M. chifengense, M. griseoflava and M. pallidorosea, were discovered in the northeast of China. Melanoleuca chifengense is morphologically characterised by its grey to yellowish-grey pileus, decurrent lamellae, grey to yellowish-brown stipe, yellowish-grey context, ellipsoid basidiospores with irregular warts and lack of hymenial cystidia. Melanoleuca griseoflava is mainly characterised by its greyish-brown pileus, adnexed to adnate lamellae, greyish-yellow context, fusiform cystidia and almost reticulate basidiospores. Melanoleuca pallidorosea is characterised by its pinkish-white pileus, white and decurrent lamellae, ellipsoid basidiospores with round and scattered warts and lack of hymenial cystidia. The phylogenetic relationship of the three species was determined by the analyses of the ITS region and the combined data matrix (ITS-nrLSU-RPB2), respectively. The results showed that the three species formed three independent lineages. Based on the combination of both morphological and molecular data, M. chifengense, M. griseoflava and M. pallidorosea were confirmed to be new species. The morphological similarities of the three new species is also discussed.
Agaricales, morphology, phylogenetic analysis, Pluteoid clade, taxonomy
Melaleuca Pat. was erected by Patouillard in 1887. As the name ‘Melaleuca’ was found to be the same as that of a plant species,
The species of Melanoleuca are often characterised by having a convex to slightly depressed pileus, mostly hymenial cystidia, amyloid ornamented basidiospores and all hyphae without clamp connections (
Although Melanoleuca has been proved to be a monophyletic group, the classification system within the genus remains controversial. Based on the colour of the pileus and the size of the carpophore,
In this paper, the authors studied three Melanoleuca species collected in north-eastern China from 2017 to 2019. Morphological observation and phylogenetic analyses confirmed that they are novel taxa in the genus Melanoleuca.
All of the fungal specimens were described and photographed in the field. Specimens were dried in an electric drier and deposited in the Fungal Herbarium of Shenyang Agricultural University (
Total genomic DNA was extracted from fresh blocks of tissue, dried with silica gel using the cetyltrimethylammonium bromide (CTAB) method (
High-quality and representative sequences of Melanoleuca in previous studies (
The GenBank accession numbers of the sequences, determined in this study, are from MW258676 to MW258689 and MW281543 to MW281548 (Table
Species | Voucher collection | Origin | GenBank accession No. | ||
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ITS | nrLSU | RPB2 | |||
Melanoleuca pallidorosea |
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Xilingole League, Inner Mongolia, China | MW258676 | MW258684 | MW281543 |
M. pallidorosea |
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Xilingole League, Inner Mongolia, China | MW258677 | MW258687 | MW281545 |
M. griseoflava |
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Fuxin City, Liaoning Province, China | MW258680 | MW258685 | MW281544 |
M. griseoflava |
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Shenyang City, Liaoning Province, China | MW258681 | – | – |
M. griseoflava |
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Shenyang City, Liaoning Province, China | MW258682 | – | – |
M. griseoflava |
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Chifeng City, Inner Mongolia, China | MW258683 | MW258686 | MW281548 |
M. chifengense |
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Chifeng City, Inner Mongolia, China | MW258678 | MW258688 | MW281546 |
M. chifengense |
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Chifeng City, Inner Mongolia, China | MW258679 | MW258689 | MW281547 |
Phylogenetic placements of the three new Melanoleuca, inferred from the combined regions (ITS-nrLSU-RPB2) using MrBayes. The lineages with new species were shown in boxes. PP ≥ 0.95 and BS ≥ 75% were indicated around the branches. Accession numbers in GenBank (ITS, nrLSU, RPB2) follow the fungal names.
The epithet refers to the species found in Chifeng City in north-eastern China.
The new species is distinguished from M. exscissa in having yellowish tinct pileus and without any type of cystidia.
Pileus 30–60 mm diam., flat at first, becoming depressed at disc when mature, margin sometimes cracking, surface glabrous, grey to yellowish-grey (4B1 to 4B2), greyish-brown (4B4 to 4B6) at centre, often darker at margin. Lamellae crowded, adnate to decurrent, white to yellowish-white (4A2), 2.5–3.0 mm broad, with lamellulae, edge entire. Stipe cylindrical, 20–35 mm long × 2–5 mm diam., central, broadened at base, solid, surface grey to yellowish-grey at first (4B1 to 4B2), becoming yellowish-brown (5D8, 5E8) with age or after touching, striate, often with whitish basal tomentum. Pileus context up to 10 mm thick near stipe attachment, thin at margin, yellowish-grey (4B2), grayish brown to yellowish brown (5D3 to 5E5) in stipe cortex, up to brown (6E7) in stipe base. Odour none, taste mild. Spore print white.
Basidiospores (90/6/2) 7.0–8.5 (9.0) × 4.0–6.2(6.5) μm, av. 7.5 × 5.2 μm, Q = 1.40–1.45(1.50), ellipsoid, hyaline, amyloid, ornamentation verruculose, with irregular warts, sometimes with ridges. Basidia (20) 23–29 (30) × (7.0) 7.5–9.0 (10.0) μm, av. 26 × 8.5 μm, clavate, 4-spored, sometimes 2-spored, subhyaline. Hymenial cystidia absent, lamella edge sterile. Hymenophoral trama 42–85 μm broad, regular with thin-walled hyphae 5.5–16.5 μm diam., hyphae not pigmented. Subhymenium poorly developed. Pileipellis a cutis of numerous repent branched hyphae, 5.5–7.5 μm wide, thin-walled. Stipitipellis hyphae 3.5–8.0 μm diam., thin-walled, hyaline. Caulocystidia absent. Clamp connections absent.
On soil or meadow outside of a forest, often on the roadside near a forest. Known from north-eastern China.
China. Inner Mongolia: Chifeng City, Linxi County, Xinlin Town, Dauran Village, alt. 1201 m, 44.07°N, 118.08°E, 22 Aug 2017, H.B. Guo (
The epithet refers to the colour of the pileus which is greyish-brown.
The new species is distinguished from M. exscissa in having adnexed to adnate lamellae and fusiform cheilocystidia.
China. Liaoning Province: Shenyang City, Tianzhu Mountain, on the soil in woods, 31 Aug 2019, X.D. Yu (holotype:
Pileus 35–60 mm diam., flat at first, then gradually depressed, margin slightly inflexed when mature, surface fibrillose, greyish-brown (4B3 to 4B5), becoming deep yellow (4C6 to 4C8) at centre. Lamellae crowded, adnexed to adnate, white, 2.5–3.0 mm broad, with lamellulae, edge entire. Stipe cylindrical, 30–50 mm long × 3–5 mm diam., central, somewhat broadened at the base, fibrous, expanded at base, solid, surface yellowish-grey to greyish at first (4B2 to 4C2), becoming yellowish-brown (5E7 to 5E8) with age, striate, with whitish basal tomentum. Pileus context up to 10 mm thick near stipe attachment, thin at margin, greyish-yellow to yellowish-grey (4B4 to 4B2), yellowish-grey (4B2) in stipe cortex, whitish in stipe base. Odour none, taste mild. Spore deposit white.
Basidiospores (234/10/8) (5.0) 6.0–7.2 (8.0) × 4.0–5.0 (6.0) μm, av. 6.5 × 4.5 μm, Q = (1.30)1.45–1.55 (1.60), ellipsoid, hyaline, amyloid, ornamentation verruculose, warts with ridges, almost reticulate. Basidia (18) 20–25 (28) × (4.0) 5.0–6.5 (7.0) μm, av. 22 × 6.0 μm, clavate, 4-spored, occasionally 2-spored, hyaline. Cheilocystidia (40) 45–55 (60) × (6.0) 8.0–12.0 (15.0) μm, fusiform, thin-walled, with encrusted crystals at apex, abundance. Pleurocystidia scattered, similar to cheilocystidia. Hymenophoral trama 90–150 μm broad, regular with thin-walled hyphae 3.0–14.0 μm diam., hyphae not pigmented, lamella edge sterile, Subhymenium poorly developed. Pileipellis a cutis of numerous repent branched hyphae 7.5–10.5 μm wide, thin-walled, pigmented with light violet. Stipitipellis hyphae 3–10.0 μm diam., smooth, thin-walled, pigmented. Caulocystidia of two types of cells, (1) 40–90 × 6.0–10.0 μm, fusiform, thin-walled, some with encrusted crystals at apex, similar to cheilocystidia; (2) 30–40 × 7.0–10.0 μm, clavate, thin-walled, without crystals. Clamp connections absent.
Solitary, saprotrophic on the soil, on the grass, on roadsides, in woods. Known from north-eastern China.
China. Liaoning Province: Shenyang City, Tianzhu Mountain, on the soil in woods, 31 Aug 2019, X.D. Yu (
The epithet refers to the species which has a pallid rose pileus.
The new species is distinguished from M. grammopodia and M. leucopoda in having a pinkish-white pileus.
China. Inner Mongolia: Xilingole League, Xiwuzhumuqin Banner, on the grass in woods, 1051 m alt., 44.48°N, 117.86°E, 22 Aug 2017, X.D. Yu (holotype:
Pileus 30–65 mm diam., flat, with depressed centre, margin slightly undulating, expanding to uplifted, sometimes slightly lacerate when mature, surface glabrous, camel (9E8 to 10E8) at centre, pinkish-white (10A3 to 10A4) towards the margin. Lamellae rather distant, adnate to decurrent, white, 3.0–4.5 mm broad, with lamellulae of two lengths, but not intervening, edge entire. Stipe cylindrical, 20–50 mm long × 5–8 mm diam., in upper part of stipe apricot (6C8 to 6D8), becoming yellowish-brown (5E8) towards base, with whitish flocculose apex, longitudinally striate, with whitish basal tomentum. Context up to 2–5 mm thick at the pileus base, whitish to creamy, whitish in stipe cortex and base. Smell fungoid smell, taste mild. Spore print white.
Basidiospores (130/7/4) (6.5) 7.0–8.5 (9.0) × 5.0–6.0 (6.5) μm, av. 7.4 × 5.5 μm, Q = (1.28)1.31–1.40(1.44), ellipsoid, hyaline, ornamentation verruculose, warts mainly round and scattered, amyloid. Basidia (20) 25–33 (35) × (6.0) 6.5–9.5 (10.5) μm, av. 28 × 8.5 μm, clavate, 4-spored, occasionally 2-spored, subhyaline. Hymenial cystidia absent. Lamella edge sterile. Hymenophoral trama 95–159 μm wide, regular, with thin-walled hyphae, 5.0–10.0 μm diam., hyphae not pigmented. Subhymenium poorly developed. Pileipellis a cutis of numerous repent branched hyphae, 4.0–10.0 μm wide, inflated cell to 21.0 μm, thin-walled. Stipitipellis hyphae 7.0–10.0 μm, thin-walled, hyaline. Caulocystidia absent. Clamp connections absent.
Solitary or in small group, saprotrophic in grass. Known from north-eastern China.
China. Inner Mongolia: Xilingole League, Xiwuzhumuqin Banner, on the grass in woods, 1051 m alt., 44.48°N, 117.86°E, 22 Aug 2017, X.D. Yu (
Line drawings of the three new Melanoleuca species A–C Melanoleuca chifengense (holotype,
Morphologically, the most distinctive features of M. pallidorosea are a pinkish-white pileus, a yellowish stipe, white and decurrent lamellae, lack of hymenial cystidia, ellipsoid basidiospores with round and scattered warts, 7.0–8.5 × 5.0–6.0 μm. According to the classification system of
Melanoleuca chifengense is easily recognised by its grey to yellowish-grey pileus, decurrent lamellae, grey to yellowish-brown stipe and yellowish-grey context, and lack of hymenial cystidia. Melanoleuca griseoflava is characterised by a greyish-brown pileus, adnexed to adnate lamellae, yellowish-grey stipe, greyish-yellow context and fusiform cystidia. The two species have similar-sized basidiomata and grey pileus, Melanoleuca griseoflava differs from M. chifengense by the adnexed to adnate gills and having abundant fusiform cystidia. According to
In the present study, both phylogenetic analyses, based on a single region (ITS) and three regions (ITS-nrLSU-RPB2), showed that there were nine clades in the genus Melanoleuca (Figures
Sincere thanks to Dr. Zhang M, Dr. Wang CQ, Dr. Sun QB for their kind help during the field trips. This study was supported by the National Natural Science Foundation of China (No. 31770014) and Science and Technology Plan Project of Liaoning Province (2020-MZLH-33).